Schistosoma mansoni cercariae exploit an elastohydrodynamic coupling to swim efficiently

Abstract

The motility of many parasites is critical for the infection process of their host, as exemplified by the transmission cycle of the blood fluke Schistosoma mansoni. In their human infectious stage, immature, submillimetre-scale forms of the parasite known as cercariae swim in freshwater and infect humans by penetrating through the skin. This infection causes Schistosomiasis, a parasitic disease that is comparable to malaria in its global socio-economic impact. Given that cercariae do not feed and hence have a finite lifetime of around 12 hours, efficient motility is crucial for the parasite's survival and transmission of Schistosomiasis. However, a first-principles understanding of how cercariae swim is lacking. Via a combined experimental, theoretical and robotics based approach, we demonstrate that cercariae propel themselves against gravity by exploiting a unique elastohydrodynamic coupling. We show that cercariae beat their tail in a periodic fashion while maintaining a fixed flexibility near their posterior and anterior ends. The flexibility in these regions allows an interaction between the fluid drag and bending resistance: an elastohydrodynamic coupling, to naturally break time-reversal symmetry and enable locomotion at small length-scales. We present a theoretical model, a 'T-swimmer', which captures the key swimming phenotype of cercariae. We further validate our results experimentally through a macro-scale robotic realization of the 'T-swimmer', explaining the unique forked-tail geometry of cercariae. Finally, we find that cercariae maintain the flexibility at their posterior and anterior ends at an optimal regime for efficient swimming, as predicted by our theoretical model. We anticipate that our work sets the ground for linking the swimming of S. mansoni cercariae to disease transmission and enables explorations of novel strategies for Schistosomiasis prevention.